Jessica Bouwmeester

Towards a phylogenetic classification of reef corals: the Indo-Pacific genera Merulina, Goniastrea and Scapophyllia (Scleractinia, Merulinidae)

Recent advances in scleractinian systematics and taxonomy have been achieved through the integration of molecular and morphological data, as well as rigorous analysis using phylogenetic methods. In this study, we continue in our pursuit of a phylogenetic classification by examining the evolutionary relationships between the closely related reef coral genera Merulina, Goniastrea, Paraclavarina and Scapophyllia (Merulinidae). In particular, we address the extreme polyphyly of Favites and Goniastrea that was discovered a decade ago. We sampled 145 specimens belonging to 16 species from a wide geographic range in the Indo‐Pacific, focusing especially on type localities, including the Red Sea, western Indian Ocean and central Pacific. Tree reconstructions based on both nuclear and mitochondrial markers reveal a novel lineage composed of three species previously placed in Favites and Goniastrea. Morphological analyses indicate that this clade, Paragoniastrea Huang, Benzoni & Budd, gen. n., has a unique combination of corallite and subcorallite features observable with scanning electron microscopy and thin sections. Molecular and morphological evidence furthermore indicates that the monotypic genus Paraclavarina is nested within Merulina, and the former is therefore synonymised.

Pachyseris inattesa sp. n. (Cnidaria, Anthozoa, Scleractinia): a new reef coral species from the Red Sea and its phylogenetic relationships

Abstract A new scleractinian coral species, Pachyseris inattesa sp. n., is described from the Red Sea. Despite a superficial resemblance with some species in the agariciid genus Leptoseris with which it has been previously confused, P. inattesa sp. n. has micro-morphological characters typical of the genus Pachyseris. This genus, once part of the Agariciidae, is comprised of five extant species and is widely distributed throughout the tropical Indo-Pacific. It is currently incertae sedis as a result of recent molecular analysis and appears to be closely related to the Euphylliidae. A molecular phylogenetic reconstruction including P. inattesa sp. n., the genus type species P. rugosa, and P. speciosa, all present in the Red Sea, was performed using the mitochondrial intergenic spacer between COI and 16S-rRNA. The results confirm that P. inattesa sp. n. is a monophyletic lineage closely related to the other Pachyseris species examined.

First biological measurements of deep-sea corals from the Red Sea

It is usually assumed that metabolic constraints restrict deep-sea corals to cold-water habitats, with ‘deep-sea’ and ‘cold-water’ corals often used as synonymous. Here we report on the first measurements of biological characters of deep-sea corals from the central Red Sea, where they occur at temperatures exceeding 20°C in highly oligotrophic and oxygen-limited waters. Low respiration rates, low calcification rates, and minimized tissue cover indicate that a reduced metabolism is one of the key adaptations to prevailing environmental conditions. We investigated four sites and encountered six species of which at least two appear to be undescribed. One species is previously reported from the Red Sea but occurs in deep cold waters outside the Red Sea raising interesting questions about presumed environmental constraints for other deep-sea corals. Our findings suggest that the present understanding of deep-sea coral persistence and resilience needs to be revisited.

Synchronous spawning of Acropora in the Red Sea

Multi-specific synchronous spawning is a reproductive strategy used by scleractinian corals that has now been described from coral reefs in 23 locations globally (Baird et al. 2009). While high multi-specific synchrony in the reproductive condition of Acropora colonies has been documented in the Red Sea in April and/or May (Hanafy et al. 2010), multi-specific synchronous spawning has not been directly observed. In April 2011, mature oocytes were found in a high proportion of colonies in numerous species of Acropora on reefs near Thuwal, Saudi Arabia (22 18¢19.26†N, 38 57¢56.66†E). On the night of April 16 2011, two nights before the full moon, egg-sperm bundles were first observed in Acropora polyps at 20:30 h (Fig. 1a). Between 22:30 and 23:45 h, 43 colonies from 10 out of 13 surveyed Acropora species released egg/sperm bundles (Fig. 1b), including three species that had not been observed to spawn previously (A. plantaginea, A. parapharaonis, and A. lamarcki). This is the first documented multi-specific synchronous spawning event in the Red Sea, demonstrating that the asynchronous spawning pattern at Eilat in the Gulf of Aqaba (Shlesinger and Loya 1985) is not representative of the Red Sea, and providing further support for the prediction that these events are characteristic of all speciose coral assemblages (Guest et al. 2005).

Forgotten in the taxonomic literature: resurrection of the scleractinian coral genus Sclerophyllia (Scleractinia, Lobophylliidae) from the Arabian Peninsula and its phylogenetic relationships

The monospecific scleractinian coral genus Sclerophyllia Klunzinger, 1879 was originally described from Al-Qusayr (Egypt) in the Red Sea based on a series of solitary specimens. Thenceforth, it has been considered a junior synonym of Symphyllia and Cynarina based on corallum macromorphology. In this study, several specimens of Sclerophyllia margariticola were collected on the coasts of Saudi Arabia in the northern and central Red Sea. Four molecular markers were sequenced, COI and the intergenic spacer between COI and l-rRNA from mitochondrial DNA and Histone H3 and ribosomal ITS2 from nuclear DNA. Phylogenetic trees and haplotype network analyses show that S. margariticola belongs to the family Lobophylliidae and that it is closely related to Acanthastrea maxima, an uncommon species from waters around the Arabian peninsula (the Gulf of Aden, Arabian Sea, Gulf of Oman and Persian Gulf). Sclerophyllia margariticola and A. maxima share several macro- and micromorphological characters, such as the presence of free septa, high elliptical septal teeth perpendicular to the septal margin, irregular lobate tips, very wide tooth spacing, a very strong granulation with granules scattered all along the septal sides, and a palisade interarea structure, and their micromorphology differs substantially from that of Acanthastrea echinata, the type species of Acanthastrea. Therefore, we formally resurrect Sclerophyllia, provide a revised diagnosis for the genus, and move A. maxima into Sclerophyllia.

First report of tubular corallites on Stylophora caused by a symbiotic copepod crustacean

Amongthemorethan340speciesofsymbioticcopepodsassociatedwithnumerous scleractinian species, four have been reported from galls orcysts in three species of corals (Dojiri 1988; Kim and Yamashiro 2007).The relationship between these copepods and their scleractinian hostsremains poorly studied. Here, we provide the first report of a tubularcorallite shape caused by copepods.The copepods were found to create short tubular outgrowths (2 mmdiameter) on Stylophora pistillata branches in the Saudi Arabian RedSea, off Thuwal (22 03¢48.5†N, 38 45¢51.2†E) on April 29, 2013(Fig. 1a). Each outgrowth is a modified thick-walled corallite with aterminal opening and a polyp inside (Fig. 1b, c). The inner chamber ofthe modified corallite has longitudinal folds atypical for Stylophora aswell as a reduced columella (Fig. 1c, d). Outgrowths found on S. pi-stillataintheRedSeadifferfromthosepreviouslyreported(Dojiri1988;Kim and Yamashiro 2007) due to the presence of a terminal openingframed by well-developed tentacles of the polyp (Fig. 1b).Single female individuals of the crustacean copepod, representing anew species of the genus Spaniomolgus Humes and Stock 1972 (Poe-cilostomatoida:Rhynchomolgidae),werefoundwithinthegastriccavityof the affected polyps (Fig. 1e). The genus Spaniomolgus was previ-ously known only from Madagascar. Nonscleractinian invertebrates areparticularly understudied in the Red Sea, and further work is needed tofully characterize their diversity and function in this region.

Cyphastrea kausti sp. n. (Cnidaria, Anthozoa, Scleractinia), a new species of reef coral from the Red Sea

Abstract A new scleractinian coral species, Cyphastrea kausti sp. n., is described from 13 specimens from the Red Sea. It is characterised by the presence of eight primary septa, unlike the other species of the genus, which have six, ten or 12 primary septa. The new species has morphological affinities with Cyphastrea microphthalma, from which it can be distinguished by the lower number of septa (on average eight instead of ten), and smaller calices and corallites. This species was observed in the northern and central Red Sea and appears to be absent from the southern Red Sea.

High reproductive synchrony of Acropora (Anthozoa: Scleractinia) in the Gulf of Aqaba, Red Sea

Coral spawning in the northern Gulf of Aqaba has been reported to be asynchronous, making it almost unique when compared to other regions in the world. Here, we document the reproductive condition of Acropora corals in early June 2014 in Dahab, in the Gulf of Aqaba, 125 km south of previous studies conducted in Eilat, Israel. Seventy-eight percent of Acropora colonies from 14 species had mature eggs, indicating that most colonies will spawn on or around the June full moon, with a very high probability of multi-species synchronous spawning. Given the proximity to Eilat, we predict that a comparable sampling protocol would detect similar levels of reproductive synchrony throughout the Gulf of Aqaba consistent with the hypothesis that high levels of spawning synchrony are a feature of all speciose coral assemblages.

Spatial variation in coral reef fish and benthic communities in the central Saudi Arabian Red Sea

Local-scale ecological information is critical as a sound basis for spatial management and conservation and as support for ongoing research in relatively unstudied areas. We conducted visual surveys of fish and benthic communities on nine reefs (3–24 km from shore) in the Thuwal area of the central Saudi Arabian Red Sea. Fish biomass increased with increasing distance from shore, but was generally low compared to reefs experiencing minimal human influence around the world. All reefs had a herbivore-dominated trophic structure and few top predators, such as sharks, jacks, or large groupers. Coral cover was considerably lower on inshore reefs, likely due to a 2010 bleaching event. Community analyses showed inshore reefs to be characterized by turf algae, slower-growing corals, lower herbivore diversity, and highly abundant turf-farming damselfishes. Offshore reefs had more planktivorous fishes, a more diverse herbivore assemblage, and faster-growing corals. All reefs appear to be impacted by overfishing, and inshore reefs seem more vulnerable to thermal bleaching. The study provides a description of the spatial variation in biomass and community structure in the central Saudi Arabian Red Sea and provides a basis for spatial prioritization and subsequent marine protected area design in Thuwal.